Download EVOLUTION - Carol Lee Lab - University of Wisconsin

EVOLUTION
The Unifying Concept in Biology
Dr. Carol Eunmi Lee
University of Wisconsin, Madison
“Nothing in biology
makes sense except in
the light of evolution”
Theodosius Dobzhansky (1900-1975)
Who am I?
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BA, MA from Stanford University
Anthropology (Human Evolution)
PhD, University of Washington
Evolutionary Genetics
Postdoc, University of California, San Diego
Evolutionary Physiology and Biochemistry
Professor, University of Wisconsin, Madison
Center of Rapid Evolution, Zoology, Genetics
TA
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Marijan Posavi: Ph.D. Animal Science, Ph.D. candidate in
Zoology
Will lead optional discussion (times posted on website)
He has office hours and can help with understanding the
course material, but, he is NOT a private tutor (not possible
for ~100+ students)
Please contact me if you are having trouble with this course
and need a tutor
Research in
my Lab
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Adaptation, Functional evolutionary
genomics, Physiological Evolution
Rapid evolution of invasive species
entering the Great Lakes (zebra
mussels, quagga mussels, copepods)
Evolution of microbial species carried
by these invaders
Course Website
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https://mywebspace.wisc.edu/carollee/web
/Lee/Evolution410.html
***Background needed for
this course
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Some understanding of basic genetics
(Hardy Weinberg Equilibrium, DNA,
RNA, transcription, translation, allele,
genotype)
Results from diagnostic quiz
30
25
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N = 98
Mean = 5.59 ± 2.00
20
18
Frequency
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15
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10
9
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1
0
0
0
1
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Score
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Evolution of a
Pathogen as an
Example:
I will now use an infectious disease to illustrate
basic evolutionary concepts.
The following example illustrates several
evolutionary mechanisms
I will explain these concepts in more detail over
the next few lectures
HIV: Fastest
evolving organism
on Earth
HIV infects
macrophages,
T-cells
AIDS Pandemic
AIDS is among the most deadly epidemics in Human
History (1981-2012: ~36 million deaths)
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~34 million people living with AIDS (estimated 2010)
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90 million deaths predicted by 2020
#people living with HIV
UNAIDS. 2008 Report on the
Global AIDS Epidemic
(http://www.unaids.org/en/Knowle
dgeCentre/HIVData/GlobalRepor
t/2008/)
A global view of HIV infection
33 million people [30–36 million] living with HIV, 2007
Problem :
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HIV has the fastest mutation rate of any virus or
organism observed to date
HIV evolves more rapidly than humans, and more
quickly than the ability of humans to produce new
drugs
Implications: AIDS vaccines are unlikely to work on all
strains of the virus… …and unlikely to work on a given
strain in the long run
Our understanding of how to combat viruses had in general been
poor, and the recent intensive research on HIV has greatly
enhanced our understanding of how to combat viruses in general
Evolutionary
History of HIV
HIV evolved multiple
times from SIV (Simian
Immunodeficiency Virus)
There are multiple strains of HIV
of different evolutionary origins
and with different properties
(coat proteins are divergent)
Time
http://www.sciencemag.org/cgi/c
ontent/full/313/5786/462
HIV
 Retrovirus with two single
strand RNA genomes
 Uses Reverse Transcriptase
to replicate RNA --> DNA
 Attacks host immune system:
infects host macrophages and
helper T cells
(Review in textbook on HIV)
HIV virion
latches onto
proteins on
surface of host
cell
Evolutionary Forces operating
during HIV infection
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Genetic Drift (Lecture 6): Viruses and bacteria in
general have huge population sizes, where genetic drift
tends to be less important
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Mutation (Lecture 7): Fastest mutation rate known.
Lots of genetic variation is produced. Virus can respond
very quickly to Natural Selection
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Natural Selection (Lecture 10):
Selection is imposed by the host immune system
 Very strong selection is imposed by drugs
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Transmission rate selects on levels of Virulence (tendency to
cause disease). When transmission of virus to new hosts is
rapid, selection favors strains that are more harmful to host
Evolution in a Host-Parasite System
(Coevolution)
I will now discuss how NATURAL SELECTION acts on HIV:
(1) Drugs impose Selection on HIV:
--> evolution of drug resistance
(2) Transmission Rate imposes Selection on HIV:
--> evolution of virulence
(3) Host immune system also imposes selection on the
virus HIV  talk more about this in lecture on Selection
(1) Selection in Response to Drugs
Several Types of Drugs:
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Reverse transcriptase (RT) inhibitors, interrupts early
stage of virus replication. Example: AZT
Protease inhibitors, interrupts virus replication at a later
step in its life cycle
These drugs do not cure people of AIDS, they only retard
the progress of the disease
Why does AZT work initially but fail in the long run?
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AZT (Azidothymidine) is a thymidine mimic which stops
reverse transcription and impedes viral replication
Mutations in the viral reverse transcriptase gene of HIV
arise, such that the enzyme can recognize AZT and cease
AZT
to incorporate it
These mutations slow down viral genome replication (as
the reverse transcriptase becomes more careful), but
confer resistance to AZT (Tradeoff between speed vs.
accuracy of reverse transcription)
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These slow and careful mutants are favored by selection
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Viral resistance to AZT evolves within human hosts… (takes
about 6 months)
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When AZT therapy stops, back mutations that restore
the Amino Acid sequence to the original state are then
favored by selection so that reverse transcription could
speed up again (fast & sloppy are favored – because fast
replicating mutants would outgrow the slower)
AZT (Azidothymidine) is a thymidine mimic which stops
reverse transcription and impedes viral replication
• In the presence of AZT, Selection
favors mutants that are resistant to
AZT
• Resulting in a change in % in the
population, toward a higher % of
AZT resistant mutants
Why does AZT work short term but fail in
the long run?
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The drug AZT imposes Selection on the virus.
High levels of genetic variation (generated by
high mutation rate) allows the virus to
respond to selection and evolve drug
resistance.
HIV has a high mutation rate because its
reverse transcriptase is extremely error
prone, causing mutations to often arise in
the viral genome.
Occasionally a mutation can be favorable (as
in the case of resistance to AZT), and would
be favored by selection, leading to an
increase in % of the new favorable mutant in
the population of viruses
AZT
(2) Selection on Virulence of HIV
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Need to keep host alive long enough to get
passed on to the next host
(Evolutionary tradeoff between fast population growth
versus keeping the host alive)
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High Transmission rate : High Virulence
(Can grow fast and jump to the next host; ok if host dies;
the genetic strain that grows faster will win)
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Low Transmission Rate : Low Virulence
(More virulent strains would die with the host and get
selected out; less virulent strain that does not kill the
host will win)
Selection on Virulence
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High Transmission: High Virulence
High Transmission Rate
If the virus is likely to move to a new host, the faster
growing (and more virulent) strain is likely to overtake
the slower strains and “win”
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It’s ok to kill the host, since the chances of jumping
to a new host is high
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Natural selection will favor the MORE virulent strain
Selection on Virulence
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Low Transmission: Low Virulence
Low Transmission Rate
If the virus is not likely to move to a new host the
slower growing (and less virulent) strain is likely to
“win”
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It’s not ok to kill the host, since the chances of
jumping to a new host are low. If the virus kills the
host, it will kill itself
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Natural selection will favor the LESS virulent strain
Evolution in Host-Parasite System
SELECTION ON THE HOST (Humans)
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Some humans have resistance to some HIV
strains. Likely that the proportion of people with
resistant alleles will increase.
Gene Therapy? Could we win an arms race?
But HIV evolves faster than we do
and more quickly than our ability to produce new drugs
The Geographic Spread of the CCR5 Δ32 HIVResistance Allele
Novembre et al 2005, PLOS Biology
Frequency of CCR5 Δ32 allele (Figure 1.20 in Freeman & Herron)
Combating HIV
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Must lower transmission rate of HIV
so that less fatal strains evolve
Must understand evolutionary
properties of a disease:
–Evolutionary history
–Mutation rate
–Selective Forces
–Evolutionary Tradeoffs
– Evolution in response to drug AZT: slow &
accurate vs. fast & sloppy replication
– Evolution in response to transmission rate:
slow growing & less virulent (keep host
alive) vs. fast growing & more virulent
Sample Exam Question:
1: Why is it difficult for scientists to develop drugs that
could cure patients of HIV? Pick the most correct answer.
(A) Because population growth rate of HIV is so high
(B) Because mutation rate of HIV is so high
(C) Because transmission rate of HIV is so Low
(D) Because selection acting on HIV is so high
Sample Exam Question:
2: Which of the following would most likely reduce the
virulence (lethality) of HIV, the virus that causes AIDS?
(A) Applying a greater diversity of antiviral drugs to the AIDS
patient
(B) The widespread use of contraceptives (such as condoms) that
would reduce the probability of transmission to the next host
(C) Increasing transmission rate of the virus such that a less virulent
strain evolves
(D) Reducing rate of mutation rate by using drugs that slow down
reverse transcriptase, such as AZT
History of
Evolutionary Thought
Dr. Carol Eunmi Lee
University of Wisconsin, Madison
Today’s OUTLINE:
(1) Development of Darwin’s Thought
(2) Lamarck vs. Darwin
(3) The Contribution of Darwin
(4) The Contribution of Mendel
(5) Conflict between “Mendelism” and “Darwinism”
 Next time, the “Evolutionary Synthesis” of Mendel
and Darwin
Charles Darwin (1809-1882)
Origin of
Species
is one of the
most influential
texts of this
century
Charles Darwin (1809-1882)
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His father was a doctor (Richard Darwin) and his
grandfather was Erasmus Darwin, also a doctor and a
prominent scholar who was already thinking about
evolution
Initially he studied Medicine at the University of Edinburgh
He neglected his studies and could not stand the sight of
surgery or the sight of blood (had nausea, fainted)
Sent by his father at Christ’s College, Cambridge University
to become an Anglican priest
He still neglected his studies, but he did pursue a
passionate interest in Natural History
He became close to botany professor Rev. John Henslow
and met other leading naturalists who saw scientific work
as religious natural theology (and viewed adaptation [i.e.
evolution] as God acting through laws of nature)
Reverend John Stevens Henslow (1796–1861)
“I fully believe a better man never walked this earth”
Charles Darwin to J.D. Hooker, 1861
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Darwin’s Botany professor at Christ’s College
(Divinity School), Cambridge University
Henslow encouraged students to make observations
of their own, rather than being spoon-fed
After Darwin completed his final examination in 1831,
Henslow persuaded him to begin studying geology, and
made arrangements for him to take Adam Sedgwick’s
geology course and a trip to map geological strata in
North Wales
Shortly thereafter, in 1831, Henslow encouraged
Darwin to join the Voyage of the Beagle as the
ship’s naturalist
More on Henslow:
Darwin's Mentor: John
Stevens Henslow
1796–1861
By Walters & Stow
Voyage of the Beagle
HMS Beagle, a ship sent by
British Navy to chart coastline of
South America
HMS Beagle off of South America
During the 5-year Voyage
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Darwin was supposed to be primarily a
geologist; He had some expertise in
geology, beetle collecting and dissecting
marine invertebrates
He made observations on geological
formations as the ship was mapping the
coastline
At Punta Alta in Patagonia he made a
major find of fossil bones of huge extinct
mammals in cliffs beside modern
seashells, indicating recent extinction with
no signs of change in climate or
catastrophe (“extinctions happen”)
The Galapagos Islands
Archipelago of volcanic
islands distributed around
the equator in the Pacific
Ocean, 972 km (525 nmi)
west of continental
Ecuador
The Galapagos Islands
Why were these islands so important in
shaping Darwin’s thinking?
The Galapagos Islands
• As an archipelago, there were many islands with different
populations of a “species” on each island
• The populations were related, but were slightly different on each
island… (not clear boundaries)
• This led Darwin to conclude that the populations descended from a
common ancestor… and he ultimately concluded that they had
evolved and were evolving…
•  led Darwin to question the stability of the concept of a “species”
Darwin’s Mockingbirds
"My attention was first thoroughly aroused by
comparing together the various specimens ... of
the mocking-thrush"
C. Darwin, The Voyage of the Beagle (1839)
• On Sept 17 1835, Darwin disembarked from the Beagle in the Galápagos
near Sappho Cove on Chatham Island (now known as Isla San Cristóbal),
and was struck by the bold, terrestrial mockingbird.
• The bird is known today as Mimus melanotis, the San Cristóbal
Mockingbird. Over the next 6 weeks that Darwin spent in the Galápagos, he
observed mockingbirds on three other islands.
• Darwin noticed the mockingbirds differed between islands. Nicolas
Lawson, acting Governor of Galápagos, told Darwin that the tortoises also
differed from island to island.
• Towards the end of the voyage, Darwin speculated that the distribution of
the mockingbirds and the tortoises might "undermine the stability of
Species”. Species might not be fixed entities, but are changing over time…
Galapagos Islands
Darwin’s Mockingbirds
(he did not focus that much on the finches; they
were studied more later by the Grants)
Tortoises and other animals
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Mockingbirds on different islands differed slightly in
size, shape and coloration
And all shared some traits (homologous traits)
He concluded that these birds must have shared a
common ancestor--and were not independently
created
Significance of
the HMS Beagle
Voyage
• Collections to study later
• Exposed Darwin to geological formations, fossils embedded in
strata – animals that no longer exist today and are clearly extinct –
this was known, but Darwin got to see this himself
• Exposure to animal diversity, related populations within species
living in different habitats:
1. “Species” are not immutable fixed entities
2. Organisms are related by common ancestry (tree-like
branching)
3. They are changing and branching in response to the
environment
Return to England
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Returned and discussed his
geological and biological finding
with other scholars, influenced
by Malthus, etc.
Married the wealthy Emma
Wedgewood
Did not have to work, and could
study the samples he collected,
perform experiments, make
observations, think, and write
books for the rest of his life
Historical Context:
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By mid-1800s scientific context was in place
for development of theory of Evolution
Developments in Geology:
Discovery that earth is more than 6,000 years old
Fossil Record showed change in species over time
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http://www.ucmp.berkeley.edu/history/evotmline.html
Influences on Darwin
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Geology: Darwin had Lyell’s Principles of Geology on
board the HMS Beagle
Lamarck ’s (1744-1829): Concept of Adaptation,
Inheritance of acquired characteristics
Malthus (1766-1834): Competition within species
and struggle for survival
Influences on Darwin
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Geology: Darwin had Lyell’s Principles of Geology on
board the HMS Beagle, given to him by captain of the
HMS Beagle (Robert FitzRoy)
Fossil record:
The earth is old
Many animals that once existed are now extinct
There are layers (strata) in the fossil record
show a pattern of change
Jean-Baptiste Lamarck
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French Naturalist (1744-1829)
“Professor of Worms and
Insects” in Paris
The first scientific theory of
evolution (inheritance of
acquired characteristics)
Influences on Darwin
Lamarck ’s (1744-1829) theory of Evolution
Inheritance of acquired characteristics
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Proposed the most influential mechanism of
evolution before Darwin:
Introduced the idea of Adaptation, though he
was mostly incorrect on the predominant mechanism
(though sometimes correct; will discuss Epigenetic Inheritance)
Influences on Darwin
Thomas Malthus (1766-1834):
“Essay on the Principle of Population”
Population
The idea of competition:
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food
Malthus’ calculations showed that rate of population
growth was greater than rate of increase in food supply
Thus, not all individuals could possibly survive
Based on this fact of competition, Darwin concluded that
individuals that are better adapted would be the ones that
survive and leave more offspring (and win the competition)
Malthus
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From Malthus, Darwin
came up with the idea of
competition and the
“struggle for survival” as
a component of Natural
Selection
The scientific community was
primed for a theory of Evolution
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Darwin was one of many scholars at his time that were
thinking about Evolution
Many scholars accepted the idea of Evolution at the time…
what puzzled them was the mechanism
In fact, Alfred Wallace (1823-1913) came up with the idea of
natural selection independently
Darwin rushed to publish Origins in 1859 when he learned of
Wallace’s work
Alfred Russel Wallace (1823-1913)
“On the Tendency of Varieties to Depart Indefinitely from
the Original Type”
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Based on his work in the
Malay Archipelago
He also studied populations
on different islands and came
to the same conclusion as
Darwin
Alfred Russel Wallace (1823-1913)
Father of Biogeography… focused on adaptation to
the environment
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In 1858 Wallace sent Darwin a copy of
his manuscript
Darwin was shocked that Wallace came
up with the idea of natural selection
independently
They decided on a joint presentation at
the Linnean Society in 1858, but it
received little attention
After which Darwin rushed to publish his
book in 1859
Charles Darwin (1809 -1882)
On the Origin of Species (1859)
 Living
species are related by
common ancestry
 Change
through time occurs at
the population not the organism
level
 The
main cause of adaptive
evolution is natural selection
Darwin’s Main Points
(1) Organisms Evolve (Darwin not first)
(2) Common Descent: species arise from common ancestors
(tree thinking) (Wallace also)
(3) Gradualism: changes are gradual (still debated)
(4) Population Speciation: change in proportions of
individuals having a trait in a population (Wallace also)
(5) Natural Selection: mechanism (Wallace also)
Lamarck
vs Darwin
Before Darwin had developed his ideas on the
mechanism of evolution, Lamarck had proposed an
alternate mechanism that was popular at the time
Lamarck
vs Darwin
(1) Different View on pattern of Evolution
Lamarck: Linear progression
 Darwin: Tree-like branching process
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(2) Different View on Mechanism of Evolution
Lamarck: inheritance of acquired traits (not mutations)-Individual evolution
 Darwin: Evolution at the Population Level via Natural
Selection
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Lamarck’s View on Pattern of
Evolution
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Continuum between physical and
biological world (followed
Aristotle)
Being
Realm of
Being
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Scala Naturae (“Ladder of Life” or
“Great Chain of Being”)
“Transmutation of Species” –one
species turns into another
Linear progression from primitive
to advanced
God
Angels
Demons
Man
Animals
Realm of
Becoming
Plants
Minerals
Non-Being
What is wrong with a ladder?
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Evolution is not linear but
branching
Living organisms are not ancestors
of one another
The ladder implies progress
Darwin envisaged Evolution as a Tree
The affinities of all the beings of the same class have
sometimes been represented by a great tree. I believe
this simile largely speaks the truth……
…The green and budding twigs may represent
existing species; and those produced during former
years may represent the long succession of extinct
species…..
….the great Tree of Life….covers the earth with
ever-branching and beautiful ramifications
Charles Darwin, On the Origin of Species; pages 131-132
The only figure in Origin of Species
Charles Darwin's 1837 sketch, his first
diagram of an evolutionary tree from his
First Notebook on Transmutation of
Species (1837) on view at the the Museum
of Natural History in Manhattan.
Interpretation of handwriting: "I think case
must be that one generation should have as
many living as now. To do this and to have
as many species in same genus (as is)
requires extinction . Thus between A + B
the immense gap of relation. C + B the
finest gradation. B+D rather greater
distinction. Thus genera would be formed.
Bearing relation" (next page begins) "to
ancient types with several extinct forms"
Lamarck’s View on Mechanism
of Evolution
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Individuals are evolving
If you got a tan, you’d pass it on
Lamarck’s View on Mechanism
of Evolution
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Individuals are evolving
If you got a tan, you’d pass it on
Some balls get tan
during their lifetime
Lamarck’s View on Mechanism
of Evolution
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Individuals are evolving
If you got a tan, you’d pass it on
Tan balls pass on their acquired trait to their offspring
(We now know that acquired traits could get passed on sometimes
when epigenetic modifications are inherited)
Darwin’s View on Mechanism of
Evolution
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Natural Selection
A heritable difference
(now known to be due
to different mutations)
Darwin’s View on Mechanism of
Evolution
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Natural Selection
Selection might
favor this mutation
Darwin’s View on Mechanism of
Evolution
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Natural Selection
Greater Fitness: individuals with
this mutation survive and leave more
offspring
Darwin’s View on Mechanism of
Evolution
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Natural Selection
The individuals themselves are not changing, but the
population is changing in composition (% of alleles)
Darwin’s contribution:
“Population Speciation as a result of Natural Selection”
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More offspring are produced than can survive
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Limited resources and competition for resources
(“The Struggle for Survival”)
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There is variation in a population
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Individuals better adapted to environment survive
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Survivors leave more offspring (“Survival of the
Fittest”)
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Thus, average character of population is altered
So… Darwin did NOT originate
the idea of Evolution…
… BUT he provided a plausible
Mechanism (along with Wallace)
… and much evidence
Chapters of Origins (1859)
On the Origin of Species by Means of
Natural Selection, or the Preservation of
Favoured Races in the Struggle for Life
1 Variation Under Domestication
2 Variation Under Nature
3 Struggle for Existence
4 Natural Selection
5 Laws of Variation
6 Difficulties on Theory
7 Instinct
8 Hybridism
9 On the Imperfection of the Geological Record
10 On the Geological Succession of Organic Beings
11 Geographical Distribution
12 Geographical Distribution continued
13 Mutual Affinities of Organic Beings: Morphology: Embryology:
Rudimentary Organs
14 Recapitulation and Conclusion
Other books by Darwin on
Evolution
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The Variation of Plants and Animals under
Domestication
The Descent of Man
The Expression of the Emotions of Man and
Animals
Full list of books by Darwin:
http://en.wikipedia.org/wiki/Charles_Darwin_bibliography
But… Darwin’s theory was not complete
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Because Darwin knew nothing about mutation, he had
no idea how variation was generated in populations
(Lecture 7)
Because Darwin knew nothing about genetics or genes,
he had no idea how variation was passed on to
offspring (Mendel & Hardy-Weinberg)
Darwin did not know about nonadaptive evolutionary
forces, such as Genetic Drift (Lecture 6)
Sample Exam Question
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Which of the following was NOT part of Darwin’s
theory of Evolution?
(1) Natural Selection
(2) Populations as the units of Evolution
(3) Mutations as sources of genetic variation
(4) Survival of those who produce more offspring
Mendel’s work held part of the key to what was
missing in Darwin’s Theory
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Mendel published in 1865…
was ignored until 1900
Presented a mechanism for
how traits got passed on
“Individuals pass alleles on to
their offspring intact”
(the idea of particulate (genes)
inheritance)
Hardy-Weinburg
Equilibrium
(Lecture 3)
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Wilhem Weinberg
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January 13, 1908
G. H. Hardy
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July 10, 1908 in Science
The Mathematical
expression of Mendel’s
Principles of Inheritance
Hardy-Weinburg
Equilibrium
(Lecture 3)
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Wilhem Weinberg
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G. H. Hardy
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January 13, 1908
July 10, 1908 in Science
Could mathematically show
expectations of Mendelian
inheritance and whether
expectations are realized in
nature
PROBLEMS!
BUT… Mendel and Darwin’s ideas
seemed Incompatible
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Mendel’s principles: dealt with particulate (discrete) traits
(e.g. yellow vs. green, wrinkled vs. smooth)
BUT, Darwin observed continuous traits (e.g. beak size,
body length)
Q: So, how would continuous traits get
passed on?
Selection vs Mutations
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Mutations discovered after 1900
Q: If mutations are arising, why need
selection... … if things are just
mutating?
Controversy between
Mutationists vs Darwinists
Mutationists (+Mendelianism)
 They thought that evolution required only
mutations and passing on of discrete traits
Darwinists
 They thought that evolution required only
Natural Selection on continuous variation
Controversy between
Mutationists vs Darwinists
Controversy persisted for ~30 years up till
the 1930s, during which little progress
was made
Problem caused by:
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Binary thinking (Black or White thinking): it’s this
or that… “if I’m right, you must be wrong”
 When in fact the two or more factors might
interact
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Inability to see overarching mechanism that could
explain a wide range of phenomena: “How could
your Hardy-Weinberg (Mendel) explain the inheritance of
5.1 cm, 5.5 cm beak length (continuous characters)?”
 When in fact, another factor might make it possible
(think multiple genes, rather than one gene)
Genetic Drift (Lecture 5)
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A concept as important as Natural Selection
But, not as prominent on people’s minds
1872 Gulick: Neutral Theory (Genetic Drift)
1921 A.C. Hagedoorn produced data to support
Neutral Theory of Genetic Drift
The Modern Synthesis
1930s ~ 1940s
The synthesis of population genetics
(role of mutation, genetic drift),
paleontology, systematics
Darwin and Mendel Reconciled!!!
(Next Lecture)
Questions:
(1) What were the intellectual influences on Darwin’s thought?
(2) What role did the discoveries from the Beagle’s voyage play
in the development of Darwin’s thought?
(3) What were Darwin’s main conclusions?
(4) How did Darwin’s thought differ from that of Lamarck?
(4) What did Darwin’s theory leave out?
(5) What was Mendel’s contribution?
(6) What were the sources of confusion between the proponents
of Darwin and Mendel’s ideas?
CONCEPTS
Evolution
Population
Natural Selection
Evolutionary Adaptation
Fitness
Mutation
Genetic Drift
Recombination
Hardy-Weinberg Equilibrium
Development of Darwin’s thought
Lamarck vs Darwin
Darwinists vs Mendelists
Evolutionary Synthesis
Sample Exam Questions
1.Which of the following is TRUE regarding Darwin’s contribution to the theory
of Evolution?
(A) Darwin formulated the first comprehensive theory incorporating all
mechanisms of evolution
(B) Darwin recognized that evolution occurs at the population level (in terms of
changes in frequency of traits within populations)
(C) Darwin proposed that selection acts on mutations and other forms of genetic
variation
(D) Darwin agreed with Larmarck’s idea of inheritance of acquired characters,
but included the concept of fitness.
(E) Darwin incorporated Mendel’s work on the particulate inheritance of alleles
2. For several decades "Darwinists" and "Mendelists" battled over
the mechanisms of evolution. Which of the following did NOT
contribute to this particular conflict?
(a) Geneticists/Evolutionary biologists did not understand that
continuous and discrete traits follow the same principle of
inheritance
(b) Geneticists/Evolutionary biologists did not understand that
natural selection acts on mutations in a population
(c) Mendel worked with discrete traits, whereas Darwin worked
with continuous traits, leading to differences in perspectives on
inheritance
(d) Darwin was unaware of the mechanism or unit of inheritance
(e) Darwin was unaware of the mechanisms of Genetic Drift
3. Darwin’s development of the theory of evolution by natural
selection was NOT influenced by which of the following?
(a) The concept of adaptation to the environment, which was
proposed by Lamarck
(b) Fossils of extinct organisms
(c) Each island of an archipelago having a population with
morphological differences from those on other islands
(d) Malthus' demography showing exponential growth of
populations
(e) Changes in frequency of discrete characters from generation to
generation